Temporal consistency in the long-term spatial distribution of macroinvertebrate drift along a stream reach

Previous studies of the spatial pattern of stream invertebrate drift have focused on spatial variation at microhabitat scales or landscape scales, or temporal variation over diel or seasonal scales. None have examined consistency in spatial variation over longer time scales (>1 year). This study examined invertebrate drift density and composition at fixed locations (terminal ends of 10 riffles) each month at day and night along a 1 km reach of a 2nd order stream over a period of nearly 2 years. Consistent differences in the density of macroinvertebrate drift between riffles over 2 years were observed. The only habitat characteristic observed to be related to invertebrate drift density was the length and size of riffles above sampling sites, with larger and longer riffles producing the highest drift densities. Consistent differences in the supply of drifting macroinvertebrates along a stream reach may have implications for the supply of colonists to substrate patches and the profitability of feeding positions for drift-feeding fish and other predators. Handling editor: D. Dudgeon     

A continuous study of the total drift of freshwater shrimps, Gammarus pulex, in a small stony stream in the English Lake District

1. The objective was to determine the major factors affecting the downstream dispersal (drift) of freshwater shrimps, Gammarus pulex. Sample replication and frequency are major problems in the quantification of drift. For the first time, these problems were avoided by sampling the whole stream continuously so that all the shrimps drifting downstream at the sampling point were caught in a net emptied at dusk and dawn in 1966, and every 3 days in 1967. 2. There was no consistent seasonal pattern in drift rates, but a high proportion of annual drift was taken in only a few samples. There was a nocturnal diel pattern of drift with peaks soon after dusk and just before dawn. A power function described the significant (P < 0.001) relationship between drift and flow, and was used to neutralise the dominant effects of flow by standardising total drift over 24 h, nocturnal drift and diurnal drift (drift per 50 m³). These were all significantly (P < 0.001) related to benthos density, but not to date, temperature, or length of the night or day. 3. The relationship between drift values and the independent variables, flow and benthos density, was well described (P < 0.001) by a multiple‐regression model. Adding temperature, date, and/or the length of the night or day did not improve model fit. Variations in flow and benthos density explained 94% of the variation in total drift over 24 h, 97% of the variation in nocturnal drift, but only 44% of the variation in diurnal drift. A power function described (P < 0.001) the relationship between total drift and the volume of water sampled over 3‐day periods in 1967. Flow explained 95% of this drift variation; it was unnecessary to add another variable such as benthos density. 4. The significance of this study is that it avoided the problems associated with the quantification of drift samples. Therefore, the conclusions are more robust than those of many previous studies. A high proportion of the annual drift losses would have been undetected by intermittent sampling. Temperature, season, night or day length had no significant effect on drift densities, and the relationship between drift and benthos densities was proportional, not density dependent. The nocturnal increase in drift could not be interpreted as an antipredator behaviour. The dominance of flow and benthos density was apparent but the quantitative relationships posed further questions, especially those related to drift distances at different velocities.     

Continuous drift samples of macroinvertebrates in a large river, the Danube in Austria

SUMMARY 1. A new method is described for sampling the drift in large rivers continuously and automatically, using equipment based on the marine, continuous plankton sampler of Hardy (1936).
2. Between April and November 1989, about 790 h were sampled continuously, equivalent to a water volume of 2937m³ and the total catch was 527 drifting macroinvertebrates, in forty-nine taxa. Twenty-five of these taxa are recorded in the drift for the first time and twenty-eight of the drifting taxa were present in benthos samples from the Danube.
3. Total drift density varied between 12 and 31 animals per 100m³ of water sampled throughout the year, with a maximum of 31 animals in May, The composition of major faunal groups showed a significant seasonal pattern, with Oligochaeta and Diptera predominant in spring, Crustacea and Insecta in summer and only Crustacea in autumn. The overall density of the macrozoobenthos from October 1986 to December 1987 was about 19360 animals m⁻² and the proportion of total benthos animals, drifting at any instant in time, ranged from 0.0026 to 0.0064%.
4. The relationship between drift density day⁻¹ and mean daily discharge was described by a power-function. Total mean drift rate of macroinvertebrates in the Danube was estimated to be 13 600 000 animals per 24 h and the mean drift distance was estimated to vary between about 4 and 31 m, dependent on the animal group and the water velocity.
5. No obvious consistent diel pattern could be established from the continuous samples, and no marked diel rhythm could be detected for Oligochaeta, Diptera larvae and Crustacea.     

Fish drift in a Danube sidearm‐system: II. Seasonal and diurnal patterns

Seasonal and diurnal patterns of larval and juvenile fish drift were investigated in the Marchfeldkanal, a man‐made side branch of the Danube River near Vienna, Austria. A clear seasonal pattern with peak densities in mid‐June was found. Species composition varied over time, showed a site specific pattern and was dominated by tubenose goby Proterorhinus marmoratus . Water temperature was the main factor responsible for the increase of drift densities until the median drift date and repeated occurrence of early larval stages in drift indicated repeated spawning for many species. Significant differences in drift densities between different time periods of the day (day, dusk, night and dawn) were found for common bream Abramis brama , barbel Barbus barbus , chub Leuciscus cephalus , tubenose goby and roach Rutilus rutilus . The highest drift rates occurred at night (2200–0400 hours), with 86% of all larvae drifting during the hours of darkness. Fish larvae of different lengths drifting at different phases of the day were found for common bream, bleak Alburnus alburnus and chub, with largest larvae drifting during dusk (chub) and day (bleak and common bream). For bleak, all gudgeon species Gobio spp., tubenose goby, roach and for all cyprinid species combined, one 2 h night sample was found to be sufficient to predict the total 24 h drift.     

Relationship between the drift of macroinvertebrates and the activity of brown trout in a small stream

Brown trout Salmo trutta were most active in a small stream at night, dusk and dawn when drift rate was highest, but correlations between hourly drift rates and the trout's activity varied substantially between individuals, between different dates for a single individual, and between different periods of the daily cycle. On some occasions, the trout were responsive to the total drift rate, either at night or during the day, and on others to the largest drifting organisms only (terrestrial organisms, adults of Ephemeroptera, Diptera and Trichoptera). The study supports the idea that trout adapt their activity pattern to the abundance of drifting prey, either as generalists towards any organism, or as specialists towards the largest ones.     

The downstream drifting of larvae of Dixa (Diptera: Dixidae) in two stony streams

SUMMARY. Larvae of Dixa , especially D. puberula Loew., were a significant component of the invertebrate drift in the Walla Brook in southwest England, the Wilfin Beck and the River Duddon in the English Lake District, and the River Estibère in the French Pyrenees. The drifting of the larvae increased markedly at night and showed a definite diel periodicity with maximum numbers usually in the early hours of the night. Seasonal peaks in the density of larvae in the drift frequently occurred in months when adults and especially pupae were present. Most drifting larvae were in the fourth and final instar, and their drift rate usually increased when they were searching for suitable pupation sites.     

Seasonal and diel patterns of invertebrate drift in different alpine stream types

1. We examined the seasonal and diel patterns of invertebrate drift in relation to seston and various habitat characteristics in two each of four different kinds of alpine streams [rhithral (snow‐fed) lake outlets, rhithral streams, kryal (glacial‐fed) lake outlets and kryal streams]. Samples were collected at four times of the day (dawn, midday, dusk and midnight) during three seasons (spring, summer and autumn). 2. Habitat characteristics differed mainly between rhithral and kryal sites, with the latter having higher discharge and turbidity, lower water temperature, and higher concentrations of ammonium, and particulate and soluble reactive phosphorus. Seasonality in habitat characteristics was most pronounced for kryal streams with autumn samples being more similar to rhithral sites. 3. The concentration of seston was lowest in the glacial‐influenced lake outlets and slightly higher in the stream sites; no seasonal or diel patterns were evident. 4. The density of drifting invertebrates averaged less than 100 m^?3 and was lowest (<10 m^?3) at three of the four kryal sites. Taxon richness and diversity were lowest at rhithral lake outlets. Chironomidae dominated the drift as well as benthic communities and <30% of benthic taxa identified were found in the drift. 5. Drifting invertebrates showed no consistent seasonal pattern. However, density tended to be highest in spring at rhithral sites and in autumn at kryal sites. No diel periodicity in drift density was found at any site and the lack of diel pattern may be a general feature of high altitude streams. 6. Glacially influenced habitat parameters were a major factor affecting drift in these alpine streams, whereas no clear differences were observed between streams and lake outlets. Our findings indicate that invertebrate drift in alpine streams is primarily influenced by abiotic factors, and therefore, substantially differs from patterns observed at lower altitude.     

Effects of trout on the diel periodicity of drifting in baetid mayflies

Some benthic invertebrates in streams make frequent, short journeys downstream in the water column (=drifting). In most streams there are larger numbers of invertebrates in the drift at night than during the day. We tested the hypothesis that nocturnal drifting is a response to avoid predation from fish that feed in the water column during the day. We surveyed diel patterns of drifting by nymphs of the mayfly Baetis coelestis in several streams containing (n=5) and lacking (n=7) populations of rainbow trout, Oncorhynchus mykiss. Drifting was more nocturnal in the presence of trout (85% of daily drift occurred at night) than in their absence (50% of daily drift occurred at night). This shift in periodicity is due to reduced daytime drifting in streams with trout, because at a given nighttime drift density, the daytime drift density of B. coelestis was lower in streams occupied by trout than in troutless streams. Large size classes of B. coelestis were underrepresented in the daytime drift in trout streams compared to nighttime drift in trout streams, and to both day and night drift in troutless streams. Differences in daytime drift density between streams with and without trout were the result of differences in mayfly drift behaviour among streams because predation rates by trout were too low to significantly reduce densities of drifting B. coelestis. We tested for rapid (over 3 days) phenotypic responses to trout presence by adding trout in cages to three of the troutless streams. Nighttime drifting was unaffected by the addition of trout, but daytime drift densities were reduced by 28% below cages containing trout relative to control cages (lacking trout) placed upstream. Drift responses were measured 15 m downstream of the cages suggesting that mayflies detected trout using chemical cues. Overall, these data support the hypothesis that infrequent daytime drifting is an avoidance response to fish that feed in the water column during the day. Avoidance is more pronounced in large individuals and is, at least partially, a phenotypic response mediated by chemical cues.     

Invertebrate drift lost to the sea during low flow conditions in a small coastal stream in Western Canada

The invertebrate drift leaving the lower end of a small coastal stream on the east coast of Vancouver Island was recorded during low flow conditions. 20,156 animals, weighing 0.96 g (dry weight), were collected in drift nets over a 5 day period in early summer. High and low drifting taxa are listed. Considerable daily variation in total captures occurred and was attributed to a few dominant taxa (primarily harpacticoid copepods, mites and chironomid larvae) exhibiting atypical drift patterns. 0.004% of the stream's invertebrate standing crop was estimated to be in the water column at any instant in time. The possible use of the outgoing animals as food for juvenile salmon in the estuary is discussed.     

Diel drift of Chironomidae in a large lowland river (Central Poland)

The diel drift patterns of Chironomidae larvae were investigated in a seventh order section of the Warta River (Central Poland) over two diel cycles during May 1989. Three nets (mesh size 400 m) were installed in a cross section of the Warta River.The estimated drift density was low, but was comparable to that calculated for other large rivers. Spatio-temporal fluctuations in abundance and composition of macroinvertebrate drift, including Chironomidae, were observed with the highest density of drifting macrobenthos recorded near the depositional bank of this river. The ratio benthosdrift indicated differing propensities for of the older instars of a given chironomid taxon to drift. Orthocladiinae larvae were the most abundant subfamily of Chironomidae in drift but not in benthos, reaching up to 73% of the total drifting chironomid larvae. More taxa but fewer individuals (about 20% of the chironomid larvae collected) belonged to the tribe Chironomini, the dominant group in benthos.A major part of chironomid drift collection may represent behavioural drift because the net mesh size used in the Warta River was insufficient to catch the earliest instars (distributional drift). Both at the family and subfamily level chironomid larvae exhibited a distinct nocturnal drift periodicity. Nocturnal periodicity was documented for the dominant species, but due to the low density of many chironomid species, it was impossible to determine their diel drift pattern. Some Chironomidae appeared to be aperiodic.     

Spatial and Temporal Variation of Macroinvertebrate Drift in Two Neotropical Streams1

The detection of spatial variation in macroinvertebrate drift depends on the spatial scale of investigation in streams of the La Selva Biological Station, Costa Rica. Drift samples were taken in a spatially nested design, with two streams, two reaches per stream, two riffles per reach, and four replicate samples per riffle. Drift showed little variation among streams, but varied significantly at the scales of reach and riffle, with variation among samples also high. In addition, sampling took place at two temporal scales: diel and at two different periods that differed in rainfall conditions. Drift diel periodicity was a clear pattern, while only density of individuals varied among sampling periods. This is the first study of macroinvertebrate drift at multiple spatial scales, despite the recognition that multi‐scale studies are essential for a more complete understanding of community patterns and processes.     

Invertebrate drift in the Dan River,Israel

The Dan river, a principal source of the Jordan River, Israel, is unusually constant in discharge (8 m³·s^–1) and water temperature (15–16 °C). The Jordan headwaters constitute the southernmost oasis of a palearctic north temperate fauna, and presumably the very constancy of the Dan contributes to its important role as a regional refuge. However, little is known of river ecology from this region. We report a twelve month study of drift, undertaken to assess diel, seasonal, and spatial patterns of the abundance of drifting invertebrates.Diel periodicity in drift was detectable but minimal. Baetidae nymphs showed a pronounced nocturnal increase, gammarid amphipods a modest, twofold increase, while dipteran larvae showed no diel variation. Seasonal variation likewise was minimal and due principally to the Baetidae, while gammarid amphipods showed no significant seasonality. The notably small diel and seasonal variation in aquatic drift in the Dan may be attributable to the extremely constant physical regime.Spatial variation was substantial. Two stations located 30 and 200 m below the karstic exsurgence of the Dan provided drift densities among the lowest reported anywhere, whereas two stations located 1 and 4.5 km downstream had more typical drift densities. A water diversion project completed halfway through the study resulted in a 50% reduction in flow at the most downstream stations, but had no discernible effect on drift.     

Diel vertical and horizontal distribution of crustacean zooplankton and young of the year fish in a sub-alpine lake: an approach based on high frequency sampling

Understanding the spatial dynamics of predators and their preyis one of the most important goals in aquatic ecology. We studiedspatial and temporal onshore–offshore distribution patternsin young of the year (YOY) Eurasian perch (Perca fluviatilis)and crustacean zooplankton (Daphnia hyalina, Cyclops prealpinus)along a transect in Lake Annecy (France). Our study representsa first attempt at coupling hydroacoustic fish survey and highfrequency zooplankton recording to assess simultaneously thelarge-scale distribution patterns of YOY fish and their zooplanktonprey over a diel cycle (day, dusk and night sampling). We hypothesizedthat the spatial distribution of zooplankton could be shapedby both anti-predator behaviour (horizontal and vertical migrations)and predation losses. Fish biomass, size structure and dietwere assessed from split-beam echosounding and net trawlingsamples, whereas crustacean abundances were estimated with asmall modified Longhurst–Hardy continuous plankton recorder.We evaluated the diel changes in the spatial distribution patternsof fish and zooplankton and determined the overlap between theirdistributions. Fish biomass was dominated by YOY perch in upperwarmer layers and salmonids (Coregonus lavaretus and Salvelinusalpinus) in the colder and oxygenated deep layers. YOY perchwere aggregated in dense schools in the epilimnion during theday and dispersed at night. Fish biomass was distributed alonga strong increasing onshore–offshore gradient at night,whereas crustacean prey showed a decreasing gradient. This onshore–offshorenegative gradient in crustacean distribution, expressed on ashorter scale during the day, shifted toward the surface watersat night. A distinct kinetic of diel vertical migration (DVM)patterns was exhibited by daphnid and cyclopoid populationsand resulted in distinct vulnerability to perch predation. Spatio-temporaldistribution of crustaceans in Lake Annecy during the diel cyclestudy was probably shaped both by predation loss to YOY perchand by anti-predator behaviour (DVM, DHM) by zooplankton. Theimplications for fine-scale studies of fish-zooplankton interactionsare discussed.     

Dynamics of Invertebrate Benthic Communities and Drift in a Regulated River of Central Spain

We document invertebrate benthic and drift dynamics in a regulated river in central Spain at two temporal scales: seasonal (for both benthos and drift) and daily (for drift). The benthic abundance of individuals and taxon richness generally increased in the summer. Drift abundance showed no seasonal or daily variation, but taxon richness of drifting individuals was higher in the spring. Both ben‐thos and drift showed clear seasonal changes in taxonomic composition. Interestingly, some benthic taxa showed their highest abundances in the spring, while others were more abundant in the summer. In contrast, most drifting taxa were more abundant in the spring. Different functional feeding groups showed different patterns of variation throughout the year, both in the benthos and the drift. Daily variations in drift were present in very few taxa and functional feeding groups, and only in some seasons. (© 2005 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Interannual variability in seasonal dynamics and species composition of drifting young-of-the-year fishes in two European lowland rivers

st young-of-the-year (YOY) fishes in two lowland rivers in the Czech Republic (Danube Basin) drifted from mid-May to mid-July and almost exclusively at night. The peak of drift density lasted c . 3 weeks and depended on water temperature (by its effect on fish spawning), but not on discharge. Peak drift densities varied between years and rivers from 80 to 1354 fish 1000 m⁻³. More than 98% of drifting fish were cyprinids. Rutilus rutilus, Rhodeus sericeus, Gobio spp., Alburnus alburnus and Abramis brama were dominate in the 22 species encountered in the River Morava. Carassius auratus gibelio and R. sericeus were the most common of 17 species in the River Kyjovka. Species composition of drifting fishes was similar among years in the Morava but varied in the Kyjovka. The species composition of drifting fishes did not differ from those of the YOY fish assemblage in nursery areas during the drift season. Relative abundances in nursery areas decreased after the drift season in species that dominated in the drift, but increased in those that avoided drift. It is suggested that drift is a regular part of the life history of many cyprinid fishes and interannual variability in density and species composition of drifting YOY fish can be explained by variable spawning success among years. Differences in the relative abundances of the YOY fish assemblage in nurseries during and after the drift season suggests that drift may cause significant mortality.     

The relation between invertebrate drift and two primary controls,discharge and benthic densities,in a large regulated river

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Diel changes of larval drift among amphidromous gobies in Japan, especially Rhinogobius brunneus

Larvae of amphidromous gobies migrate to the sea as passive drift. In order to identify the diel drift pattern (DDP) of each species, we conducted 24 hour collections in different sections ofthree rivers. The DDPs ofdifferent species at each sampling station were almost identical, but the DDP of each species varied among stations, depending on the topographical features of water-courses. Downstream migration occurred throughout the day in steep water-courses while it was nocturnal with a peak in number soon after sunset in gentle gradients in the plain and was also nocturnal with a small amount of day time drifting between the two gradients. It is suggested that the larvae drifting from the upper parts of the river in daytime halted their migration in a pool with low flow rate until twilight.     

Drift dynamics of Chironomidae larvae: I. Preliminary results and discussion of importance of mesh size and level of taxonomic identification in resolving Chironomidae diel drift patterns

Diel drift samples utilizing nets with mesh size less than 200 microns were taken in Linesville Creek, Pennsylvania, an eastern deciduous forest stream, and Inlet Run, Wyoming, an alpine snow melt stream. Identification of drifting Chironomidae larvae to lowest level taxonomic categories indicated 51 species or species group categories representing 51.95% of the total insect drift in Linesville Creek and 18 species or species group categories representing 70.47% of the total insect drift in Inlet Run. Orthocladiinae were the predominant larvae in the drift in Linesville Creek, with 19 species comprising 43.84% of the Chironomidae drift. In decreasing abundance were Chironomini (12 species, 40.36% of Chironomidae drift), Tanytarsini (10 species, 8.89%), and Tanypodinae (10 species, 6.91%). By contrast, Diamesinae were the predominant larvae in the drift in Inlet Run, (5 species, 71.43%) followed by Orthocladiinae (10 species, 27.25%), Tanytarsini (2 species, 1.20%), and Podonominae (1 species, 0.12%). Comparison of drift composition with substrate samples and/ or emergence data indicated a close relationship between relative abundance in drift and relative abundance in the benthos. Behavioral drift patterns with nocturnal peaks were seen for 3 species or species groups in Linesville Creek. Four species with diurnal drift peaks were present in Inlet Run. Analysis of the size distribution of drifting larvae indicates that a mesh size as small as 200 microns is required to resolve diel drift patterns. It is postulated that random factors greatly influence the apparent diel drift pattern of Chironomidae when nets with mesh size in excess of 400 microns are employed in drift studies. Conflicting literature reports of behavioral drift for Chironomidae may be due to differing species composition of drifting larvae and net mesh size related artifacts.     

Diel,size-differential drift patterns of three macroinvertebrate species in the lower Mississippi River,Louisiana (USA)

We explored macroinvertebrate size-differential drift in the lower Mississippi River (a 9th order system). Because this river system is highly turbid, we hypothesized that visually-dependent vertebrate predators feeding on drifting organisms would be at a disadvantage. Thus, size-differential drift should not occur. For one 24-hour period in both January and April, six drift nets were used to sample surface drift. Nets were emptied once every four hours. Individual intra-ocular distances of three macroinvertebrate species (Hydropsyche orris: Trichoptera, Hexagenia limbata: Ephemeroptera, Macrobrachium ohione: Crustacea) were measured. Percentages of size classes in the drift were determined. In both months, large individuals of H. orris and H. limbata were prevalent in the nocturnal but scarce in the diurnal drift. In January, large M. ohione drifted regardless of time. In April, large M. ohione predominated the nocturnal drift. Our results could not be attributed solely to vertebrate predator avoidance. Other mechanisms such as diel microhabitat migration and current velocity may have accounted for the results.     

Aquatic and terrestrial invertebrate drift in southern Appalachian Mountain streams: implications for trout food resources

1. We characterised aquatic and terrestrial invertebrate drift in six south‐western North Carolina streams and their implications for trout production. Streams of this region typically have low standing stock and production of trout because of low benthic productivity. However, little is known about the contribution of terrestrial invertebrates entering drift, the factors that affect these inputs (including season, diel period and riparian cover type), or the energetic contribution of drift to trout. 2. Eight sites were sampled in streams with four riparian cover types. Drift samples were collected at sunrise, midday and sunset; and in spring, early summer, late summer and autumn. The importance of drift for trout production was assessed using literature estimates of annual benthic production in the southern Appalachians, ecotrophic coefficients and food conversion efficiencies. 3. Abundance and biomass of terrestrial invertebrate inputs and drifting aquatic larvae were typically highest in spring and early summer. Aquatic larval abundances were greater than terrestrial invertebrates during these seasons and terrestrial invertebrate biomass was greater than aquatic larval biomass in the autumn. Drift rates of aquatic larval abundance and biomass were greatest at sunset. Inputs of terrestrial invertebrate biomass were greater than aquatic larvae at midday. Terrestrial invertebrate abundances were highest in streams with open canopies and streams adjacent to pasture with limited forest canopy. 4. We estimate the combination of benthic invertebrate production and terrestrial invertebrate inputs can support 3.3–18.2 g (wet weight) m⁻² year⁻¹ of trout, which is generally lower than values considered productive [10.0–30.0 g (wet weight) m⁻² year⁻¹]. 5. Our results indicate terrestrial invertebrates can be an important energy source for trout in these streams, but trout production is still low. Any management activities that attempt to increase trout production should assess trout food resources and ensure their availability.